The Behavior of <Emphasis Type="Italic">Acromyrmex crassispinus</Emphasis> (Hymenoptera: Formicidae) on Trail Bifurcations and the Influence of Ant Flow on Error Rates of Nestbound Laden Workers

Ants are ordinarily faced with a succession of bifurcations along their foraging networks. Given that there is no directionality in pheromone trails, each bifurcation is potentially an opportunity for error in the trajectory of laden workers to the nest, which could entail considerable inefficiencies in the transportation of food to the colony. Leaf-cutting ants (Atta and Acromyrmex) commonly show intense traffic and complex foraging trail systems, which make them ideal organisms to study worker behavior in trail bifurcations. The behavior of leaf-cutting ants of the genus Acromyrmex in trail bifurcations is still largely unexplored. Thus, this study aimed to assess the behavior of Acromyrmex crassispinus workers on trail bifurcations and to investigate whether differences in ant flow on foraging trails influence the error rate of nestbound laden workers at trail bifurcation. There was a negative relationship between ant flow and error rate of nestbound laden workers. Most workers walked in the central part of the foraging trails but occupied a broader area of the foraging trail when the ant flow was high. The results of this study provide valuable insight into the organization of traffic flow in A. crassispinus and its impacts on the foraging strategy of the species.     

Polydomy and the organization of foraging in a colony of the Malaysian giant ant Camponotus gigas (Hym. / Form.)

In Kinabalu National Park, Borneo we observed four colonies of the Malaysian giant ant Camponotus gigas in a primary forest. These predominantly nocturnal ants have underground nests, but forage in huge three-dimensional territories in the rain forest canopies. The colony on which our study was mainly focused had 17 nests with about 7000 foragers and occupied a territory of 0.8 ha. To improve observation and manipulation possibilities, these nests were linked at ground level by 430 m of artificial bamboo trail. A group of specialist transport worker ants carried food from `source' nests at the periphery to the central `sink' nest of the queen. Transport of food between nests started immediately after the evening exodus of the foragers. Transporter ants formed a physical subcaste among the minors and behaved according to predictions of the central-place foraging theory. Their load size was about five times that of the average forager and grew proportionally with head width. Longer distances were run by ants with greater head width and larger gross weight. Transporter ants that ran more often took heavier loads. Experiments with extra-large baits revealed that C. gigas used long-range recruitment to bring foragers from different nests to “bonanzas” at far distant places. The foraging strategy of C. gigas is based on a polydomous colony structure in combination with efficient communication, ergonomic optimization, polyethism and an effective recruitment system. Received: 16 March 1998 / Accepted: 24 August 1998     

Decay rates of attractive and repellent pheromones in an ant foraging trail network

Pharaoh’s ants (Monomorium pharaonis) use at least three types of foraging trail pheromone: a long-lasting attractive pheromone and two short-lived pheromones, one attractive and one repellent. We measured the decay rates of the behavioural response of ant workers at a trail bifurcation to trail substrate marked with either repellent or attractive short-lived pheromones. Our results show that the repellent pheromone effect lasts more than twice as long as the attractive pheromone effect (78 min versus 33 min). Although the effects of these two pheromones decay at approximately the same rate, the initial effect of the repellent pheromone on branch choice is almost twice that of the attractive pheromone (48% versus 25% above control). We hypothesise that the two pheromones have complementary but distinct roles, with the repellent pheromone specifically directing ants at bifurcations, while the attractive pheromone guides ants along the entire trail. Received 15 November 2007; revised 7 March 2008; accepted 18 March 2008.     

Estimate of complexity of behavioral patterns in ants: analysis of hunting behavior in <Emphasis Type="Italic">Myrmica rubra</Emphasis> (Hymenoptera,Formicidae) as an example

A method for estimating the complexity of behavioral patterns of ants based on the Kolmogorov complexity is considered. Behavioral sequences are presented as “texts” compressed with the KGB Archiver (v. 1.2). The elements of behavior (a total of 10) singled out from video records served as an alphabet. The comparison of “successful” and “incomplete” hunting behaviors in Myrmica rubra showed that successful hunting patterns were characterized by less complexity than “incomplete” ones. It was assumed that complete patterns had less redundancy and better predictability. The smallest complexity was revealed in complete hunting patterns of naive (laboratory reared) ants in comparison with members of a natural colony. In perspective, quantitative evaluation of complexity of behavioral patterns will help to evaluate the level of discrete variability within ant colonies.     

Seed predator deterrence by seed-carrying ants in a dyszoochorous plant, <Emphasis Type="Italic">Chamaesyce maculata</Emphasis> L. Small (Euphorbiaceae)

Seeds are often carried by omnivorous ants even if they do not carry elaiosomes. Although many seeds carried by ants are consumed, both seeds abandoned during the seed carrying and leftover seeds are consequently dispersed (dyszoochory). These non-myrmecochorous seeds do not necessarily attract ants quickly. Therefore, these seeds often seem to be exposed to the danger of consumption by pre-dispersal seed predators. We propose the hypotheses, “seed predator deterrence hypothesis” that plants may benefit from seed-carrying ants if they deter seed predators from visiting plants, and seed-carrying ants may play additional roles in plant reproductive success, besides dyszoochory by ants. To test the hypotheses, we investigated the abundance of seed-carrying ants of the species Tetramorium tsushimae Linnaeus and Pheidole noda Smith F., and of the seed predatory stinkbug, Nysius plebeius Distat, on the spotted sandmat, Chamaesyce maculata L. Small, of which the seeds have no elaiosomes but are consumed by both ants and bugs. In the field, ants and stinkbugs seldom encountered each other on the plant. The number of stinkbugs beneath the plants with ants was smaller than that beneath the plants without ants. In laboratory experiments, the number of stinkbugs on the shoot was smaller when ants were present than when they were absent. These results might support the seed predator deterrence hypothesis: the probability of seed predation by stinkbugs seems to be reduced by the ant visits on plants and/or the existence of ants beneath the plants. This study highlights a new ant–plant interaction in seed dispersal by ants.     

Effect of Trail Bifurcation Asymmetry and Pheromone Presence or Absence on Trail Choice by Lasius niger Ants

During foraging, ant workers are known to make use of multiple information sources, such as private information (personal memory) and social information (trail pheromones). Environmental effects on foraging, and how these interact with other information sources, have, however, been little studied. One environmental effect is trail bifurcation asymmetry. Ants forage on branching trail networks and must often decide which branch to take at a junction (bifurcation). This is an important decision, as finding food sources relies on making the correct choices at bifurcations. Bifurcation angle may provide important information when making this choice. We used a Y‐maze with a pivoting 90° bifurcation to study trail choice of Lasius niger foragers at varying branch asymmetries (0°, [both branches 45° from straight ahead], 30° [branches at 30° and 60° from straight ahead], 45°, 60° and 90° [one branch straight ahead, the other at 90°]). The experiment was carried out either with equal amounts of trail pheromone on both branches of the bifurcation or with pheromone present on only one branch. Our results show that with equal pheromone, trail asymmetry has a significant effect on trail choice. Ants preferentially follow the branch deviating least from straight, and this effect increases as asymmetry increases (47% at 0°, 54% at 30°, 57% at 45°, 66% at 60° and 73% at 90°). However, when pheromone is only present on one branch, the graded effect of asymmetry disappears. Overall, however, there is an effect of asymmetry as the preference of ants for the pheromone‐marked branch over the unmarked branch is reduced from 65%, when it is the less deviating branch, to 53%, when it is the more deviating branch. These results demonstrate that trail asymmetry influences ant decision‐making at bifurcations and that this information interacts with trail pheromone presence in a non‐hierarchical manner.     

Polymorphism and Division of Labor During Foraging Cycles in the Leaf-cutting Ant <Emphasis Type="Italic">Acromyrmex octospinosus</Emphasis> (Formicidae; Attini)

While division of labor within leaf-cutting ant nests has been well-characterized in the context of the collection and processing of leaf material, environmental factors such as day-night cycles and heavy rainfall limit the time during which leaf-cutting ant workers leave the nest to gather forage. Using a novel “flat panel” nest design, we studied how patterns of within-nest task performance changed when a colony of the leaf-cutting ant Acromyrmex octospinosus was and was not provided access to forage. We conducted scan samples of individuals working within the nest under both conditions and compared task allocation patterns across provisioning regimes and between workers of different sizes. When labor was compared between worker size groups, “minor” workers (head width ≤2.0 mm) and “major” workers (head width >2.0 mm) showed significantly different task performance patterns when forage was available: minors performed mostly brood-care and garden maintenance, while majors were mostly involved in the handling of freshly-cut leaf fragments. In contrast, when the colony was deprived of forage, the task performance patterns of minor and major workers converged and did not significantly differ. Marked major workers known to be foragers tended to remain idle within the nest when the colony was deprived of forage, while non-foragers of similar head width engaged in a variety of within-nest tasks, suggesting polyethism in majors may be based on factors other than size.     

Ecological explanations for successful invasion of exotic plants

The intentional introduction of exotic species can increase the level of local biodiversity, enrich people’s material lives, and bring significant social and economic benefits that are also the symbols of human progress. However, along with the frequent intercourse among countries and regions, the frequency of uncontrolled crossregional migration of species is increased and there is a lack of scientific management strategy for the intentional introduction of exotic species. Exotic species invasion, which is behind habitat fragmentation, has become the second largest threatening factor to the maintenance of the global-scale level of biological diversity. Exotic species invasion can destroy the structure of an ecosystem, disturb the economic life of a society, and do harm to human health. In this paper, the authors review some of the ecological explanations for issues such as “what causes or mechanisms have led to the successful invasion of exotic species”, including the “ideal weeds characteristics”, “biodiversity resistance hypothesis”, “enemies release hypothesis”, “evolution of increased competitive ability hypothesis”, “niche opportunity hypothesis”, and “novel weapon hypothesis”. The authors also analyze and evaluate the background and theoretical basis of the hypotheses, providing explanations for some phenomena, as well as the deficiencies of these explanations.     

Pheromone trail decay rates on different substrates in the Pharaoh's ant, Monomorium pharaonis

Abstract. Many ants use pheromone trails to organize collective foraging. This study investigated the rate at which a well‐established Pharaoh's ant, Monomorium pharaonis (L.), trail breaks down on two substrates (polycarbonate plastic, newspaper). Workers were allowed to feed on sucrose solution from a feeder 30 cm from the nest. Between the nest and the feeder, the trail had a Y‐shaped bifurcation. Initially, while recruiting to and exploiting the feeder, workers could only deposit pheromone on the branch leading to the feeder. Once the trail was established (by approximately 60 ants per min for 20 min), the ants were not allowed to reinforce the trail and were given a choice between the marked and unmarked branches. The numbers of ants choosing each branch were counted for 30 min. Initially, most went to the side on which pheromone had been deposited (80% and 70% on the plastic and paper substrates, respectively). However, this decayed to 50% within 25 min for plastic and 8 min for paper. From these data, the half‐life times of the pheromone are estimated as approximately 9 min and 3 min on plastic and paper, respectively. The results show that, for M. pharaonis, trail decay is rapid and is affected strongly by trail substrate.     

Trail system in red wood ants (<Emphasis Type="Italic">Formica rufa</Emphasis> Group) under recreation press

The influence of recreation press on the forager trail system in red wood ants was examined. The model settlement of these ants was situated in green moss-pine forest, comprising areas with a different degree of the recreation press. In 2007, trail systems of 36 anthills were mapped (a total of 136 trails). In 2009, two nests from each group at each digression stage were selected as model objects. These nests possessed close to average values of the cupola diameter (d) and the number of trophic trails (n _c ). In these 6 model nests, trail systems (a total of 33 trails) were re-mapped and the strength of columns was estimated. It had been found that the recreation press decreases the population number in families of red wood ants. Trophic trails become shorter, their structure is simplified, and the number of trees with aphid colonies decreases. When the size of the family decreases owing to irregular decrease in the population number of separate columns, columns possessing non-ramified (less “efficient”) trails disappear first. Weakening of the family decreases the mobility of the trail network; trees with aphid colonies become more important.     

To be or not to be faithful: flexible fidelity to foraging trails in the leaf‐cutting ant Acromyrmex lobicornis

1. Ants using trails to forage have to select between two alternative routes at bifurcations, using two, potentially conflicting, sources of information to make their decision: individual experience to return to a previous successful foraging site (i.e. fidelity) and ant traffic. In the field, we investigated which of these two types of information individuals of the leaf‐cutting ant Acromyrmex lobicornis Emery use to decide which foraging route to take. 2. We measured the proportion of foraging ants returning to each trail of bifurcations the following day, and for 4–7 consecutive days. We then experimentally increased ant traffic on one trail of the bifurcation by adding additional food sources to examine the effect of increased ant traffic on the decision that ants make. 3. Binomial tests showed that for 62% of the trails, ant fidelity was relatively more important than ant traffic in deciding which bifurcation to follow, suggesting the importance of previous experience. 4. When information conflict was generated by experimentally increasing ant traffic along the trail with less foraging activity, most ants relied on ant traffic to decide. However, in 33% of these bifurcations, ants were still faithful to their trail. Thus, there is some degree of flexibility in the decisions that A. lobicornis make to access food resources. 5. This flexible fidelity results in individual variation in the response of workers to different levels of ant traffic, and allows the colony to simultaneously exploit both established and recently discovered food patches, aiding efficient food gathering.     

Territoriality in the Malaysian giant ant Camponotus gigas (Hymenoptera/Formicidae)

In a 5-ha area of primary lowland rain forest in Borneo, we observed four polydomous colonies of the night-active giant ant Camponotus gigas. The nonoverlapping, three-dimensional territories in the canopy had a ground size up to 0.8 ha. C. gigas showed a distinct territorial behavior: (1) specific “barrack” nests, especially containing many majors, were situated at the borders and were established during long-term territorial conflicts; (2) trunk trails were regularly patrolled by majors that attacked alien conspecifics and some other ant species violently; and (3) sentinels, often involved in long-enduring conflicts with neighboring ant colonies, defended the borders at bridgeheads. Interspecific conflicts with sympatric Camponotus species always led to violent, “bloody” fights of all castes. Intraspecific conflicts, however, were solved by ritual fights (“front leg boxing”) of majors. C. gigas performed a de-escalation strategy to end short periods of true intraspecific “ant war” that we provoked experimentally. Artificially induced ritualized combats continued over weeks also in the absence of baits, indicating that borders may become established by long-term conflicts of attrition. We discuss the differences between ritual fights in desert and rain-forest ants and apply Lanchester's theory of battles to our findings. Received: October 17, 2000 / Accepted: May 14, 2001     

An ant’s eye view of culture: propagation of new traditions through triggering dormant behavioural patterns

We consider a previously unknown way of propagation of behavioural traditions in animal communities using hunting in ants as an example. We experimentally revealed that common litter dwelling ants Myrmica rubra effectively hunt jumping prey and the way the hunting behavioural pattern is distributed within ant colonies is rather sophisticated. Comparison of our results with those obtained on vertebrates enables us to suggest that “distributed social learning” plays an important role in spreading new traditions in animal communities: initial performances by a few carriers of an “at once and entirely” available behavioural pattern propagate this pattern among specimens which have only dormant “sketches” of it. Spread of these behaviours in populations is based on relatively simple forms of social learning such as social facilitation which underlies species’ predisposition to learn certain sequences of behavioural acts. To be triggered, carriers of dormant “sketches” of a relevant behavioural pattern should encounter performances of this pattern with sufficient frequency. We call this strategy triggering of dormant behavioural patterns. Integration of behaviour thus takes place not only at the individual level but at the population level as well.     

Birds from the Far East in Central Europe: some side-notes to Pfeifer et al. (2007)

The “reverse migration hypothesis” as the explanation for the vagrancy of Far Eastern birds in Europe is commented on. It is concluded that, as the postulated switch of 180° along a great circle line would lead the eastern or even central Siberian birds to North America rather than to Europe, the application of the “reverse migration hypothesis” to them seems untenable (which, however, does not preclude its plausibility in case of transatlantic, southern European, or even southwestern Siberian vagrants). On the other hand, patterns of appearance of the Turdus [naumanni] superspecies (Machalska et al. 1967) suggest the relationship of the phenomenon of vagrancy to post-breeding nomadic movements and/or periodical invasions of non-migrating species.     

Trail traffic flow prediction by contact frequency among individual ants

Ants build a trail that leads to a new location when they move their colony. The trail’s traffic flows smoothly, regardless of the density on the trail. To the best of our knowledge, such a phenomenon has been reported only for ant species. The trail’s capacity is known as trail traffic flow. In this paper, we propose a probabilistic model of trail traffic flow, which overcomes some inadequacies of the kinetic model previously proposed in the literature. Our model answers a question unsolved by the previous model, namely, how many worker ants form such a density-independent trail. We focus on ants’ responses to mutual contacts that involve individuals in trail formation. We propose a model in which contact frequency predicts the number of worker ants that form a trail. We verify that our model’s estimates match the empirical data that ant experts reported in the literature. In modeling and evaluation, we discuss an intelligent ant species, the house-hunting ant Temnothorax albipennis, which is popular among the ant experts.     

Simple rules result in the adaptive turning of food items to reduce drag during cooperative food transport in the ant <Emphasis Type="Italic">Pheidole oxyops</Emphasis>

Insect workers cooperate to carry out a variety of tasks. One example is cooperative transport of food items by two or more ant workers, which is important in foraging in many species. We predicted that natural selection would result in strategies that improve the performance of this task and tested this in Pheidole oxyops, a Neotropical ant in which ca. 70% of the biomass of dead insects brought back to the nest is transported cooperatively. We specifically tested the hypothesis that groups would re-orient food items to reduce drag, given that legs, wings, and other projections should affect the ease of dragging prey in different orientations. By presenting ants with artificial food items and dead cockroaches, both of which required approximately twice as much force to drag backwards as forwards, and a control which was equally easy to move in both orientations, we showed that natural groups of 3–20 food-transporting ants usually turned items that were facing backwards (72 and 83% of trials for artificial food items or cockroaches, respectively), the orientation requiring greater force, but not items facing forwards (10 and 12% of trials, respectively). Turning usually involved a single ‘steering’ ant. The key role of the ‘steering’ ant was shown by removing either the current steering ant or a randomly chosen ‘non-steering’ ant during turning. In 100% of the trials in which the steering ant was removed, turning stopped until another ant took its place. Conversely, turning stopped in only 17% of trials in which a ‘non-steering’ ant was removed. Turning is an emergent property of the system and may not have been directly selected for. Rather, turning seems to occur through a combination of pre-existing retrieval behaviour and the underlying physics of large loads. Points where the food item catches the ground can act as a fulcrum or pivot around which the item can rotate. Ants furthest from the fulcrum have more leverage and so are more likely to play a key role in turning. A simple rule relevant to individual transport of food items such as “grasp the food item and move towards the nest”, when used in the context of cooperative transport, has allowed the ants to solve a seemingly complicated problem requiring coordination.     

The ‘truck‐driver’ effect in leaf‐cutting ants: how individual load influences the walking speed of nest‐mates

The foraging behaviour of social insects is highly flexible because it depends on the interplay between individual and collective decisions. In ants that use foraging trails, high ant flow may entail traffic problems if different workers vary widely in their walking speed. Slow ants carrying extra‐large loads in the leaf‐cutting ant Atta cephalotes L. (Hymenoptera: Formicidae) are characterized as ‘highly‐laden’ ants, and their effect on delaying other laden ants is analyzed. Highly‐laden ants carry loads that are 100% larger and show a 50% greater load‐carrying capacity (i.e. load size/body size) than ‘ordinary‐laden’ ants. Field manipulations reveal that these slow ants carrying extra‐large loads can reduce the walking speed of the laden ants behind them by up to 50%. Moreover, the percentage of highly‐laden ants decreases at high ant flow. Because the delaying effect of highly‐laden ants on nest‐mates is enhanced at high traffic levels, these results suggest that load size might be adjusted to reduce the negative effect on the rate of foraging input to the colony. Several causes have been proposed to explain why leaf‐cutting ants cut and carry leaf fragments of sizes below their individual capacities. The avoidance of delay in laden nest‐mates is suggested as another novel factor related to traffic flow that also might affect load size selection The results of the presennt study illustrate how leaf‐cutting ants are able to reduce their individual carrying performance to maximize the overall colony performance.     

An agent-based model to investigate the roles of attractive and repellent pheromones in ant decision making during foraging

Pharaoh's ants organise their foraging system using three types of trail pheromone. All previous foraging models based on specific ant foraging systems have assumed that only a single attractive pheromone is used. Here we present an agent-based model based on trail choice at a trail bifurcation within the foraging trail network of a Pharaoh's ant colony which includes both attractive (positive) and repellent (negative) trail pheromones. Experiments have previously shown that Pharaoh's ants use both types of pheromone. We investigate how the repellent pheromone affects trail choice and foraging success in our simulated foraging system. We find that both the repellent and attractive pheromones have a role in trail choice, and that the repellent pheromone prevents random fluctuations which could otherwise lead to a positive feedback loop causing the colony to concentrate its foraging on the unrewarding trail. An emergent feature of the model is a high level of variability in the level of repellent pheromone on the unrewarding branch. This is caused by the repellent pheromone exerting negative feedback on its own deposition. We also investigate the dynamic situation where the location of the food is changed after foraging trails are established. We find that the repellent pheromone has a key role in enabling the colony to refocus the foraging effort to the new location. Our results show that having a repellent pheromone is adaptive, as it increases the robustness and flexibility of the colony's overall foraging response.     

Moving targets: collective decisions and flexible choices in house-hunting ants

Many decisions involve a trade-off between commitment and flexibility. We show here that the collective decisions ants make over new nest sites are sometimes sufficiently flexible that the ants can change targets even after an emigration has begun. Our findings suggest that, in this context, the ants’ procedures are such that they can sometimes avoid ‘negative information cascades’ which might lock them into a poor choice. The ants are more responsive to belated good news of a higher quality nest than they are when the nest they had initially chosen degraded to become worse than an alternative. Our study confirms, in a new way, that ant colonies can be very powerful “search engines”.     

The advantages and disadvantages of being introduced

Introduced species, those dispersed outside their natural ranges by humans, now cause almost all biological invasions, i.e., entry of organisms into habitats with negative effects on organisms already there. Knowing whether introduction tends to give organisms specific ecological advantages or disadvantages in their new habitats could help understand and control invasions. Even if no specific species traits are associated with introduction, introduced species might out-compete native ones just because the pool of introduced species is very large (“global competition hypothesis”). Especially in the case of intentional introduction, high initial propagule pressure might further increase the chance of establishment, and repeated introductions from different source populations might increase the fitness of introduced species through hybridization. Intentional introduction screens species for usefulness to humans and so might select for rapid growth and reproduction or carry species to suitable habitats, all which could promote invasiveness. However, trade offs between growth and tolerance might make introduced species vulnerable to extreme climatic events and cause some invasions to be transient (“reckless invader hypothesis”). Unintentional introduction may screen for species associated with human-disturbed habitats, and human disturbance of their new habitats may make these species more invasive. Introduction and natural long-distance dispersal both imply that species have neither undergone adaptation in their new habitats nor been adapted to by other species there. These two characteristics are the basis for many well-known hypotheses about invasion, including the “biotic resistance”, “enemy release”, “evolution of increased competitive ability” and “novel weapon” hypotheses, each of which has been shown to help explain some invasions. To the extent that biotic resistance depends upon local adaption by native species, altering selection pressures could reduce resistance and promote invasion (“local adaptation hypothesis”), and restoring natural regimes could reverse this effect.